K RA S TEV |0 Do political and economic decision-making rely on common neural substrates? Sekoul Krastev Integrated Program in Neuroscience McGill University, Montréal August 2015 A thesis submitted to McGill University in partial fulfillment of the requirements of the degree of Masters of Science © Sekoul Krastev 2015 K RA S TEV |1 TABLE OF CONTENTS ABSTRACT ....................................................................................................................... 3 RÉSUMÉ............................................................................................................................ 5 PREFACE.......................................................................................................................... 7 CHAPTER 1: INTRODUCTION .................................................................................... 8 CHAPTER 2: Do political and economic choices rely on common neural substrates? A review and fMRI meta-analysis. ................................................................................ 12 BACKGROUND.......................................................................................................... 12 METHODS .................................................................................................................. 14 Lite rature Search ..................................................................................................... 14 Are there domain general neural responses common to subjective value in economic and political contexts? ............................................................................ 14 Comparison of Gene ralize d and Political SV Correlates..................................... 15 RESULTS..................................................................................................................... 16 DISCUSSION .............................................................................................................. 23 CHAPTER 3: Are decision behaviors similar in economic and political choice?..... 27 BACKGROUND.......................................................................................................... 27 Evidence Gathering during Binary Choice ........................................................... 29 Attentional Drift Diffusion Model .......................................................................... 30 Specific Aims ............................................................................................................ 31 Predictions.................................................................................................................... 33 METHODS .................................................................................................................. 34 Subjects ..................................................................................................................... 34 Materials ................................................................................................................... 34 Eye-Tracking ............................................................................................................ 35 Procedure.................................................................................................................. 35 Data Analysis............................................................................................................ 38 RESULTS..................................................................................................................... 39 Demographic Results ............................................................................................... 39 Choice Task Results................................................................................................. 40 K RA S TEV |2 Simulation................................................................................................................. 49 DISCUSSION .............................................................................................................. 53 Alte rnative Decision Models ................................................................................... 57 Conclusion & Future Directions............................................................................. 58 CHAPTER 4: GENERAL DISCUSSION & CONCLUSION .................................... 60 ACKNOWLEDGEMENTS ........................................................................................... 64 REFERENCES................................................................................................................ 65 K RA S TEV |3 ABSTRACT The methods of cognitive neuroscience have begun to be applied to study political behavior. The neural substrates of value-based choice have already been extensively studied in economic contexts, and might provide a powerful starting point for understanding political choice. In this thesis, I present work that addresses the commonalities and distinctions between political and economic choice, within a cognitive neuroscience framework. First, a systematic literature review was undertaken to identify papers reporting neural correlates of political behavior in humans. We then asked whether the brain regions linked to subjective value in economic choice were engaged during political choice, addressing this question with a functional magnetic resonance imaging meta-analysis. This showed that only a small number of studies of political behavior have used frameworks that are comparable to those used in neuroeconomics. Further, few of the activation foci reported in these studies of political behavior fell within areas consistently found to reflect subjective value in economic studies. This raised the interesting possibility that the neural substrates of subjective value identified in economic choice paradigms may not generalize to political choice, but also highlighted the need for political choice paradigms that would allow this question to be directly tested. As a first step in this direction, in a second study we adapted a task commonly used to study information gathering in economic choice to study hypothetical voting choices. We asked whether this methodology could be applied to measure evidence gathering in voting and explored the effect of partisanship on this process. Twelve Canadian Liberal partisans and twelve non-partisans made binary choices between photographs of K RA S TEV |4 unknown political candidates in the presence and absence of party information, while choice behavior and eye movements were measured. In the absence of party information, we found that choice behavior and eye movement patterns across groups resembled those found in economic choice studies, but partisans trended toward faster choices and made significantly fewer fixations. When party information was introduced, both groups still conformed to choice behavior and eye movement patterns consistent with those seen in economic paradigms. Although party information had a substantial effect on voting behavior and eye movements in partisans, it did not completely supersede the effects of visual information and attentional modulation present throughout a trial as would have been expected if choices were being made purely based on party information. Preliminary efforts to fit an existing computational model developed in economic choice showed that partisans’ behavior was consistent with a lowered decision threshold, and party information acted to boost the initial value of the option for partisans. This work suggests that binary choice tasks used in economic studies can be applied to analyze political decisions, and provides preliminary data on the mechanisms by which partisanship influences such choices. This thesis provides a starting point for a neuroscience-informed analysis of political decision-making behaviors, and sets the stage for work to address the neural basis of these behaviors. K RA S TEV |5 RÉSUMÉ Les méthodes de la neuroscience cognitive ont commencé à être appliquées pour étudier le comportement politique. Les substrats neuronaux des choix de valeur ont déjà été largement étudié dans des contextes économiques, et pourraient fournir un point de départ pour comprendre les choix politiques. Dans ce document, je présente des travaux portant sur les points communs et les différences entre le choix politique et économique, dans le cadre de la neuroscience cognitive. Tout d'abord, une analyse systématique de la littérature a été effectuée afin d'identifier les publications rapportant des corrélats neuraux de comportements politiques chez les humains. Nous avons ensuite demandé si les régions du cerveau liées à la valeur subjective dans les choix économiques ont été engagées au cours des choix politiques. Pour répondre à cette question, nous avons effectué une méta-analyse et les résultats ont suggéré qu’un nombre relativement faible d'études de comportement politique ont rapporté des régions du cerveau qui sont comparables à ceux utilisés dans les taches neuroéconomiques. En outre, peu des régions d’activation rapportées dans ces études de comportement politique ont tombées dans les zones liées à la valeur subjective dans les études économiques. Cela soulève la possibilité que les substrats neuronaux identifiés dans les paradigmes de choix économique ne peuvent pas être généralisés aux choix politiques. Il démontre également le besoin pour des paradigmes expérimentaux qui peuvent être comparés aux recherches existantes sur les choix économiques. Dans une deuxième étude, nous avons adapté une tâche de choix binaire couramment utilisé dans l’étude des choix économiques pour étudier les choix politiques. Nous avons demandé si cette méthode peut être appliquée pour mesurer la collecte de données pendant les choix politiques et ensuite nous avons exploré l'effet de la partisannerie sur ce processus. Douze partisans libéraux canadiens et douze non-partisans ont fait des choix binaires entre des photographies de candidats K RA S TEV |6 politiques inconnus en présence et en absence d'informations du parti, tandis que les comportements de choix et les mouvements oculaires choix ont été mesurés. En absence d'information sur la partie politique des candidats, nous avons constaté que les comportements de choix et les mouvements oculaires entre les groupes ressemblaient à ceux retrouvés dans les études de choix économiques. De plus, nous avons observé une tendance vers des choix plus rapides et un plus grand nombre de fixations visuels dans le groupe partisan. Lorsque l'information du parti politique a été introduit, les deux groupes conformaient encore aux comportements de choix et les mouvements oculaires. Bien que l'information du parti a eu un effet considérable sur le comportement de choix et les mouvements oculaires des partisans, il n’a pas remplacé complètement les effets de l'information visuel ainsi que la modération attentionnelle présente tout au long du procès – quelque chose qui aurait pu être attendu si les choix étaient faits purement basée sur l'information de partie. Des efforts préliminaires pour adapter nos données à un modèle existant développé dans le contexte des choix économiques ont montré que le comportement partisan est compatible avec une barrière de décision réduite, et que l'information du parti agit pour augmenter la valeur initiale de d’une option. Ce document suggère que les tâches de choix binaires utilisés dans les études de choix économiques, ainsi que les modèles analytiques développées dans ces contextes, peuvent être généralisés aux décisions politiques. Ensuite, il fournit des données préliminaires sur les mécanismes par lesquels la partisannerie influence de tels choix. Cette dissertation fournit un point de départ pour une analyse des comportements de prise de décisions politiques informé par la méthode neuroscientifique, et ouvre la voie pour aborder la question de la base neurale de ces comportements. K RA S TEV |7 PREFACE All of the research presented in this thesis was carried out at the Montreal Neurological Institute in Montreal, Quebec, under the supervision of Dr. Lesley K. Fellows with advice on the political science elements from Dr. Dietlind Stolle and Dr. Elisabeth Gidengil, Dept. of Political Science. The literature review presented in Chapter 2 was carried out by Sekoul Krastev and the fMRI meta-analysis was carried out by Sekoul Krastev with the help of Dr. Joe Kable and, Dr. Joe McGuire at the University of Pennsylvania. The task in Chapter 3 was adapted from work at Dr. Antonio Rangel’s laboratory at the California Institute of Technology with the guidance of Dr. Ian Krajbich. The political choice task and trial generation scripts were adapted by Sekoul Krastev from work on an aesthetic choice task created by Avinash Vaidya. Subject recruitment was done by Sekoul Krastev in collaboration with Dr. Dietlind Stolle and Dr. Elisabeth Gidengil, using a database from the Center for the Study of Democratic Citizenship at McGill University. Additional subjects were recruited from the community with the help of Christine Déry. K RA S TEV |8 CHAPTER 1: INTRODUCTION Political behavior is a complex form of social interaction, ubiquitous across human societies and ecologically instrumental for outcome generation within groups (Schreiber, 2004). Although political behavior has traditionally been studied at the group level, it emerges from decisions taken by individuals (Sniderman et al., 1993). An increasing focus on the individual in political science has given rise to the field of political psychology, which seeks to explore the role of factors such as emotion, personality, socialization, group association and conflict in political behaviours (Cottam et al., 2010). As political theories have increasingly benefited from psychological models of behavior, researchers have begun to take an interest in their biological underpinnings (Lieberman et al., 2003; Oxley et al., 2008; McDermott, 2009). During the last decade, a small body of research has emerged, using the methods of cognitive neuroscience to study political behavior. Since the primary goal of political behavior is the distribution of decision-making power to members of a group, it is, by definition, a decision behavior. The neural substrates of decision-making have been extensively studied in economic contexts and might provide a powerful starting point for investigating the biological basis of political choice. This effort, sometimes termed “neuroeconomics”, has been deeply influenced by the same neoclassical economic models of utility that have shaped discussions of political decision-making within rational choice theories of political behavior. To the extent that political choices also involve weighing options with the goal of maximizing subjective value (utility), this neuroeconomic framework may also be relevant for studying the brain K RA S TEV |9 basis of political decision-making contexts (Fellows, 2004; Kable & Glimcher, 2009; Rangel et al., 2008; Padoa-Schioppa, 2011; Rangel & Clithero, 2013). As in economic contexts, deliberative political choice is a cognitively taxing undertaking (Downs, 1957; Fiorina, 1981): in principle, a decision-maker attempts to collect and process all currently available information in order to rank alternatives based on their expected utility. Decisions are thought to involve preference formation relying on an online comparison of the values assigned to competing options (Padoa-Schioppa, 2011). While several models have been developed to explain how this happens in the brain, largely based on economic paradigms, it seems likely that multiple computational steps are involved: option identification, value assignment, action selection, outcome evaluation and learning (Fellows, 2004). The assignment of subjective value to options lies at the conceptual center of the decision-making process in this type of model and has been the focus of much of the decision neuroscience work to date (Fellows, 2004; Kable & Glimcher, 2009; Rangel et al., 2008; Padoa-Schioppa, 2011). While deliberative decision-making may have normative appeal, the motivation and cognitive work required often eclipse the amount of effort most citizens are willing to invest in political affairs (Fiske & Taylor, 1991). In an alternative model, decisionmaking can instead be driven by the use of ‘heuristics’ or cognitive shortcuts (Lupia et al., 2000). In this case, the decision process may consist of little more than the identification of alternatives and the use of simple rules of thumb to make a choice: for example, using a candidate’s party affiliation to infer issue positions (Rahn, 1993). A similar version of this deliberative – heuristic dichotomy categorizes political decision-making strategies in terms of the degree of cognitive effort involved in K R A S T E V | 10 collecting and processing relevant information. This model proposes a distinction between memory-based information processing and impression driven on-line processing in making political judgments (Lodge et al., 1989). In the former, decisions are made by recalling from long-term memory all of the available considerations that surround an issue or candidate. By weighing each consideration against the other, a rough balance of positive and negative aspects can be computed that serves as a summary judgment. For impression driven on-line processing, on the other hand, new information is judged as positive or negative upon contact and then integrated into an existing ‘judgment tally’ that summarizes previous encounters with the object under consideration. Political scientists disagree on which processes underlie political decision-making. Neuroscience evidence may be helpful here as different decision processes likely rely on distinct neural circuits. The present thesis takes initial steps in identifying overlaps between the conceptual, behavioral and neural substrates of economic and political decision-making. In Chapter 2, I present the results of a systematic review of the neuropolitics literature to date, with the results of the relevant subset of fMRI studies of political choice summarized quantitatively in relation to regions commonly associated with subjective value signaling in economic choice. In Chapter 3, I report a behavioral experiment in which I adapt a binary choice task widely used to study information gathering in economic choice contexts to study political choice. I compare choice and eye movement behaviors in this laboratory voting paradigm with patterns reported in economic versions of this task, ask whether these patterns can be mimicked by a model of attentionmediated evidence gathering, and explore the impact of providing political party K R A S T E V | 11 information to Liberal partisan and non-partisan participants. The work presented in this thesis sets the stage for studies of the neural basis of political decision-making. I provide evidence as to whether political and economic decisions rely on common processes, and a starting point for a more comprehensive brain-based understanding of decision-making that is able to accommodate decisions in both contexts. K R A S T E V | 12 CHAPTER 2: Do political and economic choices rely on common neural substrates? A review and fMRI meta-analysis. BACKGROUND Political choice is multifaceted and likely relies on a variety of decision-making mechanisms. However, in most cases a political choice is likely one where some sort of value is being computed. Thus, it would be useful to know to what extent the neuroeconomics framework is applicable to the study of political decision-making. Current neuroeconomic models propose that the assignment of value to alternatives likely involves a set of variables that represent internal (e.g. hunger) and external (e.g. cost) states relevant to the consequences of each option. There is converging evidence that the orbitofrontal cortex (OFC), ventromedial prefrontal cortex (vmPFC), and associated ventral striatum are involved in assigning subjective value to alternatives. Functional neuroimaging studies in humans have shown value-related signals in vmPFC and ventral striatum in a wide range of paradigms (Bartra et al., 2013), ventromedial frontal lesions in humans disrupt even simple value based preferences (Fellows, 2007; Henri- Bhargava et al., 2012), and electrophysiological studies in monkeys show that activity of neurons in the OFC reflect changes in stimulus value (reviewed in Padoa-Schioppa (2011). The same brain substrates seem to be involved in valuing a range of primary and secondary rewards (e.g. money, odors, food, pleasurable music, attractive faces, and social rewards). For example, Lin and colleagues (2012) found that social and monetary decisions are associated with fMRI activation in overlapping regions of human vmPFC. Similarly, Watson and Platt (2012) have found that OFC neurons encode both social and non-social rewards in non-human primates. Pegors and colleagues (2014) found common K R A S T E V | 13 activation of the vmPFC in response to both place and face attractiveness, with certain areas of the vmPFC responding only to place attractiveness and others only to face attractiveness. These findings suggest that value across many stimuli may be encoded, at least in part, in a “common currency” in the vmPFC. Consistent with this claim, a recent meta-analysis of 206 fMRI studies found a consistent set of regions where activation was related to subjective value (SV) across a range of paradigms, involving primary or secondary rewards, in social, economic, and aesthetic contexts (Bartra et al, 2013). Existing findings regarding the neural substrates of value-based choice are a strong potential starting point for the study of political choice. If, as the above presented results suggest, the same neural circuitry is involved in computing value across different modalities, then it is possible that these regions are also engaged in political decisionmaking. This possibility leads us to ask the following questions: 1. To what extent has a neuroeconomic framework been used to study the neural substrates of political choice? 2. Are the same neural regions which are engaged during economic decisionmaking tasks also engaged during comparable political choice tasks? As a first step in addressing these questions, we systematically reviewed the neuropolitics literature, asking which existing studies used designs that were sufficiently similar to the neuroeconomics literature to allow comparison. The results of fMRI studies that met those criteria were summarized using a quantitative meta-analysis to provide preliminary insights into whether there are common activation patterns related to subjective value across these two literatures. K R A S T E V | 14 METHODS Literature Search A systematic literature search was conducted to identify papers discussing the neural correlates of political behavior in humans. The search terms “politics”, “political”, “democrat”, “republican”, “brain” and “neuroscience” were used on Google Scholar and Web of Knowledge. The database searches were supplemented by manual review of the citations in these papers. Papers were included in the first stage of the literature review if their central focus was the link between political behavior and the brain, whether review articles or experimental studies. This yielded twenty-seven papers using various techniques to study the brain correlates of a range of processes including face judgment in political contexts, partisanship, motivated reasoning, political interest, political attitudes and automatic processing of political preference. Are there domain general neural responses common to subjective value in economic and political contexts? In a second step, we applied the same inclusion and exclusion criteria as a recent meta-analysis of 206 fMRI studies investigating value-based choice (Bartra et al., 2013) to this body of neuropolitics research. The reference meta-analysis included studies containing the keywords “fMRI” and “reward” and identified brain regions where activity was consistently related to behavioral measures of positive and negative SV across a wide variety of value-based decision-making tasks; none of the neuropolitics papers identified in the present search were included in that meta-analysis. Subjective value (SV), defined as the “common-currency” value attributed to available alternatives, was either directly measured through ratings or preference judgments or, in the case of monetary rewards, inferred a priori as being higher for larger amounts of money. K R A S T E V | 15 The same criteria applied to our sample of neuropolitics literature yielded English-language papers in which BOLD signal was measured with fMRI, as a function of positive and/or negative SV. As in Bartra et al. (2013), we did not limit ourselves to studies that used particular tasks or stimuli, instead accepting any experimental design that yielded a clear behavioral measure of SV, such as voting for a candidate based on a photograph of his/her face (positive SV) or rating a policy negatively on a visual analog scale (negative SV). Only experiments that used whole brain analyses to report peak activation foci in stereotactic spatial coordinates (Talairach or MNI space) and linked them to either positive or negative SV measures were included. The results of the fMRI studies that met these criteria were summarized in a whole-brain meta-analysis. Talairach coordinates were converted to MNI space and activation foci were coded according to whether they corresponded to positive or negative SV. The list of coordinates so-identified is those voxels that showed significantly increased BOLD signal in relation to a behavioral measure of either higher or lower subjective value in a political task. A map of gray-matter probability (pGM) values (derived from the ICBM Tissue Probabilistic Atlases; http://www.loni.ucla.edu/ICBM) was generated to test whether there was a consistent pattern of activation related to positive or negative SV in political decision-making tasks across this set of studies, with the null hypothesis that foci were distributed randomly in the brain. Comparison of Generalized and Political SV Correlates Next, we tested whether activation foci from the neuropolitics studies fell within the regions previously identified as consistently relating to SV in economic paradigms K R A S T E V | 16 (Bartra et al., 2013). An ROI was established for positive and negative SV based on the published meta-analysis. A mask was then generated in FSL (Smith et al., 2004); the coordinates that remained represented the overlap between the political SV foci and the SV regions reported in the Bartra et al. meta-analysis. For visualization purposes, a 5 mm radius sphere was centered on each coordinate passed through the masking phase. Since the effects in the Bartra et al. study did not show lateralization, and laterality was not tested in the source studies for political SV, data were collapsed across hemispheres. RESULTS The initial literature search yielded 27 papers (Table 1) that corresponded to our search terms and inclusion criteria. Importantly, none of the studies contained the keyword “reward” that was present in all 200 of the Bartra et al. studies. Of these 27 papers, ten were reviews, six focused on political attitudes and emotion, four on face judgment, three on party identity, three on automatic processing, two on political interest and two on motivated reasoning. Overall, this analysis shows that neuropolitics is still an emerging field with relatively few original research reports to date (Fowler and Schreiber 2008). Table 1: Neuropolitics literature identified by search Author Year Jost & Amodio 2012 Kanai et al. 2011 Oxley et al. 2008 Dawes & Fowler 2009 Title Political ideology as Motivated Social Cognition: Behavioral and Neuroscientific Evidence Political Orientations Are Correlated with Brain Structure in Young Adults Political Attitudes Vary with Physiological Traits Partisanship, Voting, and the Dopamine D2 Receptor Gene Topic Partisanship Motivated Reasoning Partisanship - Party ID Attitude Partisanship Interest Method Structural MRI Structural MRI Physiological Response Genetics K R A S T E V | 17 Implicit and Explicit Evaluation: fMRI Correlates of Valence, Emotional Intensity, and Control in the Processing of Attitudes Interest in Politics Modulates Neural Activity in the Amygdala and Ventral Striatum Us versus Them: Political Attitudes and Party Affiliation Influence Neural Response to Faces of Presidential Candidates Neural Correlates of Attitude Change Following Positive and Negative Advertisements Cunningham et al. 2004 Gozzi et al. 2010 Kaplan et al. 2007 Kato et al. 2009 Knutson et al. 2006 Politics on the brain: An fMRI Investigation Face Judgment Rule et al. 2010 Voting Behavior Is Reflected In Amygdala Response across Cultures Face Judgment Schreiber et al. 2013 Spezio et al. 2009 Tusche et al. 2013 Westen et al. 2006 Zamboni et al. 2009 Amodio et al. 2007 Dhont et al. 2011 Fowler & Schreiber Friend & Thayer 2008 2011 Red Brain, Blue Brain: Evaluative Processes Differ in Democrats and Republicans A neural basis for the effect of candidate appearance on election outcomes Automatic processing of political preferences in the human brain Neural Bases of Motivated Reasoning: An fMRI Study of Emotional Constraints on Partisan Political Judgment in the 2004 U.S. Presidential Election Individualism, Conservatism, And Radicalism as Criteria for Processing Political Beliefs: A Parametric fMRI Study Neurocognitive correlates of liberalism and conservatism A Step into the Anarchist’s Mind: Examining Political Attitudes And Ideology Through Event-Related Brain Potentials Biology, Politics, and the Emerging Science of Human Nature Brain Imaging and Political Behavior: A Survey Is Political Cognition Like Riding a Bicycle? How Cognitive Neuroscience Can Inform Research on Political Thinking Emotions in Politics Automatic processing Partisanship Interest fMRI fMRI fMRI Face Judgment fMRI Attitude Partisanship - Party ID Face Judgment Automatic processing Partisanship Motivated Reasoning fMRI fMRI fMRI fMRI fMRI fMRI fMRI Attitude Partisanship - Party ID EEG EEG Attitude Review Review Commentary Commentary Commentary Lieberman et al. 2003 Marcus 2000 Marcus et al. 1998 Linking Neuroscience to Political Intolerance and Political Judgment Review McDermott 2009 The Case for Increasing Dialogue between Political Science and Neuroscience Review Schreiber 2004 Political Cognition as Social Cognition: Are We All Political Sophisticates? Review Review Review Commentary Commentary Commentary Commentary K R A S T E V | 18 Spezio & Adolphs 2007 Theodoridis & Nelson 2012 Tingley 2006 Emotional Processing and Political Judgment: Toward Integrating Political Psychology and Neuroeconomics Of BOLD Claims and Excessive Fears: A Call for Caution and Patience Regarding Political Neuroscience Neurological Imaging as Evidence in Political Science: A Review, Critique, and Guiding Science Commentary Review Commentary Review Commentary Review Seven of these 27 studies met the inclusion and exclusion criteria applied in the Bartra et al. meta-analysis (Table 2). Of the remaining 20 studies, ten were reviews and commentaries on neuropolitics as a field. Of the remaining ten studies reporting primary results, four used fMRI, two used structural MRI, two used EEG, one used genetics and one used participants' physiological response to study various political behaviors. Interestingly, the structural MRI, EEG and genetics studies reported correlates of subjects' political characteristics rather than studying online decision behavior. Thus, even though 7/11 fMRI studies passed the Bartra criteria and were therefore judged to be using a neuroeconomic framework in the study of political choice, in the broader scope of all neuropolitics studies reporting primary results, this ratio drops to 7/17. The seven studies that passed our criteria reported data from a total of 187 subjects. Three of these seven studies investigated face judgment, while the remaining four studied motivated reasoning, political interest, attitude change in response to advertising and automatic processing of political preference. Across these seven studies, reporting either a binary contrast or a continuous parametric analysis in a total of 13 tasks, four reported regions linked to behavioral measures of positive SV (for example, positively rating a politician) and four reported regions related to behavioral measures of negative SV (for example, negatively rating a politician) (Table 2). K R A S T E V | 19 Within these seven neuropolitics studies, the whole-brain analysis of above chance clustering of foci across all studies reporting positive SV foci and all studies reporting negative SV foci from the neuropolitics studies did not yield any significant overlap across either the positive or negative SV group. The maximum overlap at a single location (5mm radius around a reported coordinate) was 75% (three out of four studies) reporting correlates of positive SV and 50% (two out of four studies) reporting correlates of negative SV. These results are not different than what would be expected by chance, although due to the small sample-size only a 4/4 match would have sufficient power to exceed chance. Figure 1 shows all foci of activation related to SV in political contexts overlaid on the ROIs identified in the Bartra et al. review. K R A S T E V | 20 Table 2: Neuropolitics studies reporting BOLD signal in relation to behavioral measures of positive or negative subjective value. SV Vale nce Positive Sample Topic Task Partisanship Agree vs. disagree with political and Political opinions in interested vs. Inte rest uninterested subjects Automatic Post-hoc ratings of politicians that Proce ssing are shown while subject is of Face s engaged in a distractor task Year Author Size Country Economic SV RO I al. Partisan USA Putamen Non- German Medial T emporal Partisan y Lobe, Caudate Non- USA, 28 Partisans Japan None 40 Partisans USA None Partisans USA Insula 25 Tusche 2013 Face et al. 20 Rule et Judgment Voting for a political candidate Political Changed preference for politician Attitude after Positive Political Ad 2010 al. Subjects Non- Gozzi et 2010 O verlaps with Kato et 2009 al. Democrats and Republicans Face viewing candidate from opposing Judgment party versus one from their own Kaplan 2007 et al. 20 Insula, Dorsal Ne gative Face Not voting for a political Judgment candidate Partisanship Viewing information threatening - Motivated to a political candidate from Re asoning subject's own party versus neutral Political Changed preference for politician Attitude after Negative Political Ad Non- Spezio 2009 et al. 24 Anterior Cingulate, Partisans USA T halamus Partisans USA None Partisans USA None Westen 2006 et al. 30 Kato et 2009 al. 40 Relatively few neuropolitics studies have used designs that can be compared to fMRI research on decision-making or value in economic contexts. Where comparison was possible, few foci from the political studies fell within areas that Bartra et al. previously identified as commonly reflecting either positive or negative SV in economic contexts. K R A S T E V | 21 Of all the foci reported (130 coordinates in total), only five passed the masking stage of our ROI analysis, i.e. fell within the brain regions consistently related to SV in rewardbased decision- making paradigms. The foci associated with positive SV are shown in Figure 1, and were reported by Gozzi et al. (2010) and Tusche et al. (2013). Gozzi and colleagues (2010) used a task that measured agreement with political opinions in groups of subjects varying in level of political interest. Agreement with political opinions in politically interested versus uninterested subjects was considered a measure of positive SV, and was related to activation in the putamen. In Tusche et al., (2013), subjects were shown photographs of familiar politicians while engaged in a distractor task. Viewing politicians previously rated more positively was related to activation in the medial temporal lobe and caudate. The foci associated with negative SV are shown in Figure 1, reported in studies by Kaplan et al. (2007) and Spezio et al. (2009). Kaplan and colleagues (2007) studied partisans looking at photographs of politicians of an opposing party versus their own party. Spezio et al., 2009 asked subjects to vote for unfamiliar politicians based on head and shoulders photographs. Both studies found significant BOLD activation in the insula that was related to viewing a political candidate that the subject disliked. The coordinates in these two studies did not overlap, however. Collapsing the data across hemispheres yielded two more foci associated with negative SV falling within the Bartra ROI, in the dorsal anterior cingulate and thalamus, both from the Spezio et al. (2009) study. All the foci overlapping with Bartra et al. (2013) ROI’s for positive and negative SV are shown in Table 2. K R A S T E V | 22 Figure 1: Foci derived from neuropolitics studies mapped onto ROI’s derived from Bartra et al. (2013). A: Positive SV Overlap in Striatum and MTL; B: Negative SV overlap in Insula, Dorsal, Anterior Cingulate and Thalamus K R A S T E V | 23 DISCUSSION A systematic review of the cognitive neuroscience literature studying political behavior found that a minority of studies address political decision-making online in a way comparable to past neuroeconomics research. Ten of the seventeen studies we found aimed to link relatively static political characteristics such as partisanship to neural variables measured with fMRI, structural MRI, EEG or genetics. The remaining seven, all of which used task-based fMRI to study political decision-making, were the focus of our meta-analysis, as they provided evidence of the neural correlates of value-based political judgments. Neuroscience research on value-based decision-making now constitutes a relatively large body of work, yielding detailed models of value-based decision-making (Fellows, 2004; Rangel et al., 2008; Kable & Glimcher, 2009; Padoa-Schioppa, 2011, amongst others). This work has found consistent activation of several brain regions related to subjective value in a range of economic tasks (reviewed in Bartra et al., 2013), arguing that these regions support domain-general value-related processes. There is also some evidence from human lesion studies that damage to the ventromedial frontal region, including vmPFC and medial OFC, at least, impairs value-based choices across a range of (largely ‘economic’) contexts (Fellows & Farah, 2007; Camille et al., 2011; HenriBhargava et al., 2012), supporting the claim that this region is necessary for value-based decisions, broadly defined. Testing the limits of this claim has implications both for brain-based models of decision-making, and for our understanding of the commonalities and differences between economic and political choice. K R A S T E V | 24 Given this, it is surprising that relatively few neuropolitics studies make use of designs or analytic frameworks analogous to neuroeconomic studies where choice is studied in a reward-processing context. The handful of studies identified in this review represents too small a sample for strong interpretations of the quantitative meta-analysis results. However, even at a qualitative level there were very few consistent foci associated with SV in political contexts. The tasks which did report foci overlapping with ROI’s from Bartra et al. (2013) were generally focused on ratings of politicians rather than more abstract concepts such as political attitudes. However, even within contrasts taken from these tasks, a relatively small fraction of foci overlapped with previously identified correlates of SV. While preliminary, this is a challenge to the view that value processing in the brain is content-general, i.e. that the same circuits are engaged regardless of the nature of the decision. Although no significant effects were detectable in the meta-analysis presented here, qualitative assessment of the few neuropolitics studies that do report foci within the ‘economic’ SV-related ROIs may be useful for hypothesis generation. In the study by Gozzi et al. (2010), political interest was used to differentiate groups of subjects that evaluated political statements. The behavioral measure of positive SV in this study was the contrast between agreement and disagreement with a statement in high versus low political interest individuals. Since economic SV experiments often require that a subject is explicitly motivated to receive a reward of the type offered during the study (e.g. food study subjects are told to fast before the session, and receive a chosen food item at the end of the experiment), it seems possible that a certain amount of motivation, such as that measured by political interest, is important for political value processes to engage neural K R A S T E V | 25 circuits common to those implicated in economic value. In the Tusche et al. (2013) study, the behavioral measure of positive SV was a positive post-hoc rating of photos of politicians viewed by subjects while they were engaged in a distractor task. In this case, results suggest that automatic processing of political preference may elicit increased activity in areas of the brain associated with economic value assignment. In the negative group, the two studies by Kaplan et al. (2007) and Spezio et al. (2009) both yielded overlapping foci from tasks which measured BOLD correlates of negative SV. In the Kaplan study, SV was measured by asking participants to look at photographs of politicians from an opposing party. In the Spezio study, subjects were asked to vote for unfamiliar politicians based on a photograph of their face. In both cases, activity in the insula was correlated with viewing the photograph of a disliked politician (as measured by party preference or vote, depending on the study). Given the insula’s previous association with emotional processing (Wright et al., 2004; Phan et al., 2002) and the fact that subjects are viewing a disliked politician, it is possible that these tasks elicit a negative emotional response. In the case of the Kaplan study, the response may be an instance of pre-established distaste for both the politician (assuming he/she is familiar) and the associated party. These preliminary findings raise the possibility that the neural substrates of SV identified in economic choice paradigms may not generalize to the types of political choices studied in neuropolitics work to date. One potential difference between political and economic choice is the extent to which decision-makers rely on heuristics. Although economic frameworks encompass choices made based on simplifying heuristics, such as brand loyalty, neuroeconomic studies have largely steered away from what political K R A S T E V | 26 science would term heuristic choices (e.g. Hauser, 2011), instead focusing on paradigms where choices more clearly hinge on deliberations about subjective value. While such a focus obviously has merits, the study of heuristic decision-making has continued to grow in prominence in political science and behavioral economics, as social science comes to better understand how humans grapple with informational complexity. Indeed, as profound ignorance of many facets of political life has become commonplace in the general public (Delli Carpini & Keeter, 1995), heuristics may serve as something of the last refuge for democratic decision-making and governmental accountability (Lupia, 1994; Popkin, 1991). Based on the prevalence of heuristics in political choice and the possible differences between the neural substrates of economic and political choice, one conclusion we can draw is that comprehensive decision neuroscience must consider choices beyond the economic realm, with more work needed to address choices where deliberative economic models of value assignment fall short. Another possibility is that there is a misalignment in the frameworks used to study these economic neuropolitics and study political designs choices. that allow Stronger ready conclusions comparison would with necessitate the existing neuroeconomics literature. We propose that such studies would be of high interest to both neuropolitics and neuroeconomics research, regardless of their outcomes. Defining the extent to which current models of subjective value apply or do not apply in political choice will enrich our understanding of the brain basis of both of these important aspects of human behavior. K R A S T E V | 27 CHAPTER 3: Are decision behaviors similar in economic and political choice? BACKGROUND In Chapter 1, we examined the extent to which the methods of cognitive neuroscience have been used in the study of political behavior. Neuropolitics is still a very young field. Some of the work to date does address political decision-making, but varies in how closely it follows frameworks emerging from neuroeconomics research. Interestingly, those fMRI studies that could be directly compared showed little overlap in the subjective value-related activations. This raises the possibility that political choice is inherently different from the ‘economic’ choices that have been the main focus of neuroeconomics to date. There are potentially many differences between political and economic choices. For one, as previously discussed, the higher informational complexity inherent to most political choices may push political decision makers to use cognitive shortcuts (Schaffner & Strebb, 2002). An obvious example is the partisanship heuristic, which allows choices to be simplified to the point of following pre-established rules (e.g. vote for the candidate representing ‘my’ party). Although some economic choices can also be simplified in this way (e.g. brand loyalty), existing neuroeconomics research has typically avoided paradigms where decisions can be made by the use of heuristics, focusing instead on deliberative choice. Overall, there is little evidence one way or the other that addresses whether political and economic decision-making relies on common processes and neural substrates. K R A S T E V | 28 We hypothesize that existing neuroeconomics models can be usefully applied to understand political choice. A first step in testing this hypothesis is to examine whether the simplifying heuristics that are an important feature of political choice can be accommodated by existing behavioral models of decision-making. Binary choice preference tasks are widely used paradigms in decision psychology and neuroeconomics. They are simple and ecologically valid, and can provide data on the relationship between prior item ratings and choice, as well as decision time. Further insight into the decision process can be gained by measuring eye movements, an established proxy for information gathering during deliberation in these tasks. Formal computational models of such tasks exist that reproduce many of the observed behavioral phenomena, providing an explicit explanatory framework that has been linked, at least provisionally, to underlying brain networks (Lim et al., 2011). Past work on evidence gathering during economic choice has found consistent patterns in the choice behavior and eye movements of subjects during value-based binary choice tasks (Milosavljevic et al., 2010). Firstly, past studies of binary economic choice have shown that the probability of choosing a given option increases as the difference between the value rating (made a priori) of that option and its alternative increases. Secondly, past studies show that reaction time (RT) decreases as the difference in value rating between the two alternatives increases. Thus, a priori value ratings are a good predictor of both choice probability and RT during binary choice. These findings have been interpreted by some as support for the idea that deliberative decisions are driven by an evidence-gathering process in the brain (Krajbich et al., 2010). K R A S T E V | 29 Visual fixation patterns from similar studies also support this idea. Visual fixations during each trial have been studied with respect to the time of stimulus presentation and choice: binned into first (the initial eye movement when the stimuli are presented), middle (all subsequent shifts in direction of gaze) and last (the fixation ongoing at the time the choice is registered). It has been shown that during economic choice, final fixations are significantly shorter than middle fixations. This has been interpreted as support for evidence gathering to a decision threshold, which, when reached, interrupts the final fixation. Furthermore, the item fixated during the last fixation is far more likely to be chosen than its alternative. One explanation for this pattern is that the value of the unattended (i.e. non-fixated) item is discounted. Thus, studies so far support the idea that deliberative choice involves an evidence gathering process that can be indexed by visual fixations. Evidence Gathering during Binary Choice Information gathering has been studied extensively in the context of perceptual decision-making, yielding several models that attempt to bridge the gap between stimulus and response (Gold & Shadlen, 2007). These models, which have largely focused on binary decisions, consider choice as a three step process: gathering evidence for each alternative; integrating random noise into the evidence; and making a choice once a sufficient level of evidence (signal:noise) has been reached that favors one alternative over the other (Bogacz et al., 2006). The Drift Diffusion Model (DDM) (Ratcliff, 1978), derived from sequential analysis and signal detection theory, is perhaps the most widely-used model in studies of perceptual processing. Human and animal studies (Gold & Shadlen, 2007) have shown K R A S T E V | 30 that the DDM provides accurate predictions of behavioral and neural metrics during binary perceptual choice (Milosvljevic et al., 2010). Attentional Drift Diffusion Model In an effort to extend this literature beyond perceptual choice to value-based decision-making, a derivative of the DDM was developed which uses the same evidence gathering principles, with eye fixations as a proxy for visual attention (Krajbich et al., 2010). This model, called the attentional Drift Diffusion Model (aDDM, see Figure 2), updates a decision value toward one of two decision barriers (representing the two alternatives being weighed) but does so faster toward the alternative that is currently being visually fixated (Figure 1). The attentional Drift Diffusion Model has been shown to predict RT, choice probability and visual fixation patterns for food stimuli (photos), within certain decision contexts such as high and low time-pressure (Milosavljevic et al., 2010), and purchasing decisions (Krajbich et al., 2012) and has been adapted to three-alternative choices (Krajbich & Rangel, 2011). Furthermore, an fMRI study using this paradigm (food stimuli) has shown that rating differences between two competing items are correlated with levels of activity in the ventromedial prefrontal cortex and striatum (Lim et al., 2011). Finally, a recent study of moral decision-making showed that the aDDM can fit decision behavior beyond strictly ‘economic’ contexts (Parnamets et al., 2014). Figure 1: attentional Drift Diffusion Model - Equation. Value at time t is equal to the value at t-1 plus d (the drift rate) times the difference between the rating previously given to the attended option (in this case the left) minus theta (the discount rate) times the unattended option (in this case the right) plus epsilon (Gaussian noise). K R A S T E V | 31 Figure 2: Attentional Drift Diffusion Model – Random Walk. The light and dark grey sections represent times during which the subject is looking left and right. The resulting drift toward the attended item is a result of the discounting of the unattended item (Krajbich et al., 2010). The aDDM’s solid grounding in perceptual choice work, its use in the study of an increasingly wide variety of economic choices, and the existing evidence relating its parameters to brain processes make it a good starting point for studying political choice in a way that can be compared to existing neuroeconomic work. Specific Aims The study presented here takes the first steps toward using the methods of cognitive neuroscience to directly compare evidence gathering during economic and political choice, beginning with a behavioral task paired with eye-tracking. Since past work on evidence gathering during economic choice has yielded specific predictable choice behavior and eye-movement patterns, we asked whether these same patterns would be apparent during political choice. In particular, we aimed to K R A S T E V | 32 1. Determine whether evidence gathering during political choice yields the same behavior and eye movement patterns as previously reported for economic choice. 2. Explore the effects of partisanship on evidence gathering as indexed by choice behavior and eye movements. To address these aims, we adapted the procedure developed in binary foodchoice research by Krajbich and colleagues (2010) and applied it to voting decisions between photographs of political candidates. Past research has shown that votes based solely on appearance can predict election outcomes, suggesting that this political choice laboratory paradigm may have some real world validity (Todorov et al., 2005). We predicted that in the absence of any information except candidate appearance, these political choices would engage very similar mechanisms to economic choices, resulting in behavioral and eye-tracking patterns similar to those reported in the existing food-choice studies. In a second step, we provided party information about some of the candidates, and compared partisan and non-partisan participants to look for behavioral and eye-tracking evidence of the use of the partisanship heuristic. We predicted that Liberal Party information would completely dominate the decisionmaking process of Liberal partisans, causing their choice behavior and eye movements to deviate from the patterns seen in prior economic paradigms in trials where they are given such information. Finally, we explored whether the aDDM captures the eye tracking behaviors observed in this political paradigm, in the presence and absence of party information. This work is a starting point in determining whether existing brainbased models of value-based decision- making can be applied to political choices. K R A S T E V | 33 Predictions When no party information is presented, we predict that partisan and non-partisan participants will exhibit comparable choice behaviors, and that these will be similar to what has previously been observed during economic choice: i.e. probability of choosing an option will increase and RT will decrease as value rating difference increases. When party information is given, we predict that these behavioural patterns will be unaffected in non-partisans, but that partisans will become insensitive to the value ratings they have previously provided, both in their choices and RT. Furthermore, we predict that partisans will exhibit faster RTs in this condition compared to non-partisans, because they will use a “choose my party” heuristic. When no party information is presented, we predict that both groups will base their decision on an evidence gathering process that has previously been observed in economic choice studies, exhibiting the same eye movement patterns described above – shorter final versus middle fixations and a higher likelihood of choosing the last seen item. However, when party information is introduced, we predict that partisans will simply explore the screen until the Liberal candidate is found. We therefore expect to see fewer middle fixations. The prediction for final fixations is less clear: both a heuristic and deliberative strategy would yield final fixations shorter than middle fixations, although possibly due different mechanisms (i.e. in the former, by triggering a stimulusrule mapping such as “choose red frame”, in the latter when the decision-value barrier for “I prefer this option” is crossed). We expect partisans in the party information condition to show no bias toward the last attended item. That is, they will choose the Liberal candidate whether it is the last one they view or not. Any subtle attention-mediated bias otherwise detectable in this task will be entirely superseded by party information. K R A S T E V | 34 METHODS Subjects Subjects were recruited from databases of healthy volunteers maintained by the McGill Centre for the Study of Democratic Citizenship and the McGill Cognitive Neuroscience Research Registry, originally recruited either through web surveys or community advertisement. All subjects were screened using a standard medical questionnaire and those with a history of significant neurological or mental health issues and/or psychoactive medication use were excluded from the experiment. They were given a political attitude and knowledge questionnaire (see Fig. 3) and assigned to one of two groups based on their answers to the questions “Which political party do you ally yourself most strongly with?” and “How strongly?” Subjects who indicated that they were “fairly strong” or “very strong” Liberals were assigned to the (Liberal) partisan group and subjects who indicated either “don’t know” (7/12 subjects) or “none of these” to the first question were assigned to the non-partisan (i.e. non-Liberal-partisan) group. Subjects were also asked about their level of political interest. They provided written informed consent and received 15$/hour to compensate them for their time and inconvenience. The research protocol was approved by the MNI Research Ethics Board. Materials The stimulus set consisted of 214 head and shoulders photographs of real-life gubernatorial candidates from American elections spanning 1995-2004. The black and white photographs were converted to bitmap format and re-scaled to a width of 200 pixels. They were presented on a 17 inch Viewsonic color monitor set to a resolution of 640 x 480 pixels and located at a fixed distance of 59 cm from the subject’s eyes. K R A S T E V | 35 Stimuli thus subtended 12.5 visual degrees. Colored frames were added to the photographs, matched for contrast with the background . Figure 3: Political Questionnaire used to establish subjects’ political interest and party affiliation. Liberal partisans were defined as those chose “Liberal” in Q2A and “Fairly Strongly” or “Very Strongly” in Q2B. Non-partisans were those who chose “None of These” or “Don’t Know” in Q2A. Eye-Tracking Visual fixations were recorded using an EyeLink 1000 desktop-mounted eye tracker to track the dominant eye of each subject. The eye-tracker was calibrated to an accuracy of +/- 1.5 visual degrees. Procedure Subjects were given on-screen instructions and told that they will make choices regarding a series of politicians. The experiment was then divided into two phases: the rating phase and the choice phase. K R A S T E V | 36 During the rating phase, subjects were asked to rate all 214 stimuli, presented in a random order, on: “How much would you want to vote for this person in a real life federal election?” (scale -3 to +3). Each photograph was presented in the middle of the screen, without any response time-limit. Participants responded by mouse-clicking on the rating scale (Fig. 4). Figure 4: Phase 1 - Rating. Photographs were rated one-by-one, without any party information. (Actual photograph not shown to respect copyright.). Trials for the next phase were then generated using a subset of the initial photographs according to the following rules: only photographs rated 0 to 3 were included; 68 (34 unknown - unknown and 34 unknown – Liberal) pairs with an absolute difference of 0 were generated; 68 (34 unknown - unknown and 34 unknown – Liberal) with an absolute difference of 1 were generated; 68 (34 unknown - unknown and 34 unknown – Liberal) pairs with an absolute difference of 2 were generated (see Figure 5). unknown party was indicated by a purple frame and Liberal party was indicated by a red frame around the candidate photographs. In order to assure that choices were not K R A S T E V | 37 biased by the salience of the frame, the frame colors were chosen to have equal contrast against the background. Figure 5: Choice stimuli selection Positively rated photographs from the rating phase were randomly assigned a purple or red frame. 198 pairs were then created, so that there would be 33 pairs with a rating difference of 0, 1, 2 in the unknown-unknown Condition and 33 pairs with a rating difference of 0, 1, 2 in the unknownLiberal condition. During the binary choice phase, the 198 pairs generated at the end of the rating phase were shown in random order. Before the trials, subjects were asked “Which politician would you vote for in a real-life federal election?” (See Figure 6). The eyetracker was used during this phase in order to record visual fixations. Prior to the presentation of each binary choice stimulus pair, the subject was required to maintain fixation on a cross at the center of the screen for 2 seconds. Subjects were then presented with the two stimuli and asked to indicate their choice by pressing one of two K R A S T E V | 38 keys, without a time limit. Following completion of this phase, subjects were debriefed – they had the chance to ask questions about the general purpose of the study and were asked about their partisanship and level of political interest to confirm the intake screening. Figure 6: Choice phase. The pairs generated from Phase 1 were shown in Phase 2, and subjects were asked to choose between them, based on how likely they would be to vote for them in a real-life federal election. Data Analysis Initial data collection was accomplished on E-Prime 2.0 and EyeLink, yielding a time-series of eye fixation coordinates. Raw data were then processed with a Matlab script that identified whether each fixation location fell within an ROI (i.e. within the bounds of one of the two stimuli). An uninterrupted sequence of time-points during which fixations fell within the bounds of one stimulus was counted as a single ‘fixation’ on that stimulus, following the original methodology (Krajbich et al., 2010) (terminology that we note differs from the convention in the oculomotor literature). K R A S T E V | 39 The choice and eye-tracking data were analyzed using mixed ANOVAs, followed by Simple Effects tests and post-hoc Tukey’s HSD. Error bars on the graphs presented below indicate the standard error of the mean. RESULTS Demographic Results Forty-eight people were screened with the political questionnaire, and the 40 who met criteria as either non-partisans or Liberal partisans were included in the study. The study design required a minimum number of positively rated items in the rating phase in order to generate the binary choice phase. Twelve subjects were excluded after the rating task for not reaching this threshold – this is a relatively high number compared to past studies on economic choice, likely related to the fact that previous experiments used images of primary rewards (i.e. food), while the photographs of unknown (actual) political candidates used here were not especially appealing in the absence of other information. A further 4 subjects were excluded because the eye-tracker could not consistently follow their gaze even after several recalibrations (likely due to eye color, glasses and eye size). Full data are available for the remaining 24 subjects. Demographic and political interest data for these subjects is shown in Table 1. The Liberal partisan and non-partisan groups did not differ significantly in age, sex, education or political interest (t-tests, all p > 0.05). N Females Age Education (y) Political Interest (0-2) NonPartisans 12 5 48.8 (9.8) 15.9 (1.6) 1.5 (0.5) Partisans 12 7 45.1 (11.3) 15.5 (2.5) 1.3 (0.6) Table 1: Demographic information and level of political interest of the sample, split by extent of (Liberal) partisanship, expressed as counts or mean (SD). K R A S T E V | 40 Choice Task Results Choice We first examined basic performance on the choice task. There was no significant bias toward choosing the option presented on the left or right in either group: a mean of 50% (SD 5%) and 52% (SD 5%) of choices were for the left option in partisans and nonpartisans, respectively. As expected, a priori value ratings predicted choices. In the no party information condition, a mixed ANOVA yielded a significant main effect of rating on choice (F(4, 88) = 55.05, p < 0.01) but no significant main effect of party on choice nor a significant party-rating interaction (Figure 7a). In the party information condition, a mixed ANOVA once again showed a significant main effect of rating on choice (F(4, 88) = 15.59, p < 0.01) as well as a significant interaction between partisanship and rating (F(4, 88) = 7.14, p < 0.01) (Figure 7b). Note that in this condition, new information is available that was not used to make the initial rating, causing us to predict that partisans would no longer be affected by rating. Indeed, a simple effects test showed a significant effect of rating on choice in non-partisans (F(4, 88) = 21.45, p < 0.01), but not in partisans. However, there is evidence that the partisans were not exclusively following a simple ‘pick my party’ heuristic: when party information was present, partisans chose the Liberal candidate only 87% (SD 12%) of the time. RT RT differed significantly between groups (mean (SD): partisans 1897 (795) ms; non-partisans 3530 (2350) ms). We tested whether RT related to decision ‘difficulty’ as K R A S T E V | 41 operationalized by rating difference between candidate pairs. In the no party information condition, a mixed ANOVA yielded significant main effect of rating on RT (F(2, 44) = 4.43, p < 0.05) and a trend towards an effect of group on RT (F(1, 22) = 3.04, p = 0.09), but no interaction between group and rating (Figure 7c). In the party information condition, a mixed ANOVA also yielded a significant main effect of rating on RT (F(2, 44) = 3.77, p < 0.05) and of group on RT (F(1, 22) = 5.02, p < 0.05), but no significant interaction (Figure 7d). Tukey’s HSD revealed a significant difference between ratings 1 and 2 in non-partisans (p < 0.05), but in none of the partisan rating contrasts, suggesting that in this condition non-partisans are sensitive and partisans are insensitive to ratings. To follow up the unexpected finding that partisans trended towards faster choices than non-partisans, we examined the pattern of decision time across trials. A time-series reveals that the difference is apparent immediately, and persists throughout (Figure 8). K R A S T E V | 42 Figure 7: Choice behavior results showing choice probability and RT across groups and conditions. K R A S T E V | 43 Reaction Time Across Trials 12000 8000 6000 4000 2000 0 1 5 9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89 93 97 101 105 109 113 117 121 125 129 133 137 141 145 149 153 157 161 165 169 173 177 181 185 189 193 197 Reaction Time (ms) 10000 Trial Number Figure 8: Reaction times across all trials, showing that reaction time differences are present from the beginning of each session and thus suggesting that they are likely not an artifact of the task design. K R A S T E V | 44 Eye Movement Results Next, we examined fixation patterns. As expected, we found no significant bias toward the right or left (mean (SD): partisans 8(79) ms; non-partisans 69 (124) ms). We also found that a majority of final fixations (mean (SD): partisans 75% (12%); non-partisans 74% (10%)) were toward the item that was ultimately chosen, as in prior work. Final vs Middle Fixation Length We then tested the prediction that final fixations would be shorter than middle fixations. In the no party information condition, a mixed ANOVA yielded a main effect of fixation type on fixation duration (F(1, 22) = 15.89, p < 0.01), and a trend toward an effect of party fixation duration (F(1, 22) = 3.41, p = 0.08), but no interaction between group and fixation type (Figure 9). Tukey’s HSD revealed that final fixations were significantly shorter than middle fixations for both non-partisans (p < 0.05) and partisans (p = 0.05). Although fixation duration did not differ significantly between groups, a one-tailed t-test revealed that the number of fixations per trial was significantly lower in partisans (mean (SD): partisans: 3.0 (0.8); non-partisans: 3.7 (1.3); p < 0.01). In the party information condition, a mixed ANOVA yielded a significant main effect of fixation type on fixation duration (F(1, 22) = 19.96, p < 0.01), and a trend toward an effect of party on fixation duration (F(1, 22) = 3.64, p = 0.07), but no interaction between group and fixation type (Figure 9). Tukey’s HSD revealed that final fixations were significantly shorter than middle fixations for both partisans (p < 0.05) and non-partisans (p < 0.01). Although fixation duration did not differ significantly between groups, a one-tailed t-test revealed that the number of fixations was significantly lower in partisans (p < 0.01; mean (SD): partisans: 2.4 (0.9); non-partisans: 3.5 (1.2)). K R A S T E V | 45 Probability of Choosing Last Seen Item We next tested whether there was a choice probability advantage for the last seen item. In the no party information condition, a mixed ANOVA yielded a significant main effect of rating on choice (F(4, 88) = 32.00, p < 0.01) but no significant effect of partisanship or interaction between partisanship and rating (Figure 10a). Importantly, at rating difference 0, the last attended item had a 74% (SD 12%) chance of being chosen in both groups, which differs significantly from what we would expect if there was no advantage for the last attended item (50%), confirming the pattern seen in studies of economic choice. In the party information condition, due to the assumed rating boost provided by identifying a candidate as Liberal to a Liberal partisan, the analysis considered instances where the last attended item was the Liberal candidate separately from instances where it was the unknown-party candidate. When the unknown-party candidate was attended to last, a mixed ANOVA yielded a significant main effect of group on the probability of choosing the last seen item (F(1, 22) = 10.26, p < 0.01) as well as a significant main effect of rating on choice (F(4, 88) = 15.41, p <0.01), but no significant interaction between group and rating (Figure 10b). At rating difference 0, the last attended item was chosen by non-partisans 64% (11%) of the time and by partisans 25% (9%) of the time, showing the different impact of party information in the two groups . However, when the last attended unknown candidate was rated 1 or 2 points higher than the Liberal alternative, partisans chose that candidate 52% (SD: 13%) and 49% (SD: 11%) of the time, suggesting that party information was being integrated into the information gathering process, rather than triggering a completely separate heuristic strategy “blind” to information other than party affiliation. K R A S T E V | 46 When the Liberal candidate was the last attended, a mixed ANOVA yielded a significant main effect of group on the probability of choosing the last seen item (F(1, 22) = 17.71, p < 0.01) as well as of rating on choice (F(4, 88) = 12.31, p < 0.01). Partisans were far more likely to choose the Liberal candidate (mean (SD) at rating difference 0: 96% (7%) for partisans; 85% (9%) for non-partisans). There was also a significant interaction between partisanship and rating difference (F(4, 88) = 6.59, p < 0.01) (Figure 10c). Simple effects tests showed a significant difference between groups at rating differences -2 (F(1, 87) = 31.36, p < 0.01) and -1 (F(1, 87) = 8.53, p < 0.01). Rating also had a significant effect on the probability of choosing the last seen item in non-partisans (F(4, 88) = 18.41, p < 0.01) but had no significant effect in partisans. K R A S T E V | 47 Fixation Length First Fixation 1200 Middle Fixation Final Fixation Fixation Legnth (ms) 1000 800 600 400 200 0 No Party Info Party Info NON PARTISAN No Party Info Party Info PARTISAN Figure 9: First, Middle and Final fixation lengths are compared across groups and conditions, showing significantly shorter final versus middle fixations across all groups and conditions. K R A S T E V | 48 Figure 10: Choice probability curves for the last attended item across a) no party information trials, b) party information trials where the last attended is the unknown candidate and c) party information trials where the last item attended is the Liberal candidate. K R A S T E V | 49 Simulation The aDDM model was initialized with published parameters that capture behavior in food choice versions of this task (drift rate of 0.0002; discount rate of 0.3; Gaussian noise with sigma 0.02; barriers of +/-1; initial decision value of 0), and the parameters were then systematically varied to explore which most closely mimicked the observed group and condition differences. A time-series simulation was run with 1000 cycles and a rating scale of -2 to 2, varying parameters at 10% increments. We hypothesized that one of drift rate, discount rate, noise or barrier change would explain group differences in the No Party information trials, as these parameters affect both choice alternatives symmetrically. Similarly, we hypothesized that varying the initial decision value would explain condition differences in the Liberal partisans, as this is the only parameter which, as in this condition, can affect the two alternatives asymmetrically (i.e. favoring the one to which party information is added). Further, we expected the initial value would have changed, had the a priori ratings been made with party information provided. Parameter Fitting The default parameter values were used in the first simulation (Fig. 11a), yielding a choice probability curve that was identical to the one observed in the no party information condition (all simulated data points fell within the 95% confidence interval of the observed data (+/- 0.083)). However, while the RT simulation yielded a curve that was similar in shape to the observed data, it lay between the observed non-partisan and partisan curves: The simulated RT’s were slower than those seen in partisans and faster than those seen in non-partisans. We therefore asked which parameter change would yield a curve falling within the 95% confidence interval of the RT data we observed. K R A S T E V | 50 The only parameter change that yielded data consistent with the observed RT curve in the partisan no party information condition was lowering the decision threshold from 1 to 0.9 (see Figure 11b). A simple downward transposition fit the simulation curve to the data. In contrast, increasing the drift rate affected the slope of the curve in a way that deviated from the observed data (Figure 11c). Similarly, increasing the Gaussian noise sigma value affected both the RT and choice probability curves, making both much flatter than observed (Figure 11d). Finally, changing the starting decision value to 0.7 flattened both the choice probability curve and the RT curve, and shifter the latter downward (Figure 11e). These changes approximated the data in the partisans (party information) condition, with 4/5 choice probability points and 5/5 RT points falling within the 95% confidence interval of the data. Thus, new relevant information provided to partisans can be modelled by a shift in the starting decision value of the aDDM. This finding fits the common sense expectation; it is likely that value ratings for individual candidates would be higher in Liberal partisans if party information had been provided when the ratings were made. The preliminary model fitting suggests that group differences can be captured by changes in the decision barrier in the aDDM. While further experimental and modelling work is needed to make stronger interpretations of this finding, it raises the possibility that partisans and nonpartisans differ on the level of evidence required to make a political decision. K R A S T E V | 51 K R A S T E V | 52 Figure 11: Matlab simulation results using 1000 runs of the aDDM equation with a) default parameters, b) lowered decision barrier, c) increased drift rate, d) increased noise and e) a shifted starting decision value K R A S T E V | 53 DISCUSSION The first aim of this study was to determine to what extent political choice conforms to behavioral patterns previously observed in economic choice, when studied in a similar paradigm. The condition in which no party information was provided is the closest to existing economic work: participants had only the appearance of each candidate to guide their voting choice, and hence engaged in what was presumably on online computation of value. We found that choice probability and RT patterns in non-partisans corresponded to what has been reported in past studies with non-political stimuli, with the probability of choosing a particular candidate increasing as rating difference increased, and RT decreasing as rating difference increased. However, the patterns of behavior were different in the Liberal partisan group. Unexpectedly, there was a trend for differences in partisan and non-partisan choice behavior even in the ‘no party information’ condition. Although partisans were as likely as non-partisans to choose the candidate they had rated higher, their RT’s tended to be faster and they made significantly fewer fixations even in the absence of party information. Since this condition was informationally equivalent for the two groups, our findings indicate that either partisans in general, or Liberal partisans specifically, tend to gather less information (assuming that RT and fixations are indicative of evidencegathering) than non-partisans before making a choice. Preliminary fitting of aDDM parameters suggests that this might reflect a lower decision threshold amongst partisans. Past work lends some support to this idea (Carney et al., 2008), characterizing Liberals as being more openminded and novelty-seeking (as assessed by standard personality measures) compared to conservatives, but to our knowledge, there has been no work comparing them to non-partisans, nor showing that they exhibit faster RT’s. Further studies are needed to determine whether this K R A S T E V | 54 apparent RT difference can be replicated, to test whether it holds for partisans of other parties, whether it is present in non-political decision tasks, and whether it reflects a general personality feature or is specific to political choice. The task and analytic approach developed here would be suited to addressing these questions, with the inclusion of an economic choice condition, such as between food stimuli as studied in other work (e.g. Krajbich et al., 2010). A perceptual control condition involving a sensory rather than value-based decision would address whether the faster choices of partisans reflect a general lowering of response threshold. . Finally, standard personality questionnaires, similar to those used by Carney and colleagues (2008) to identify open-mindedness and novelty seeking in liberals could also address whether partisanship is one facet of a more general personality style, or specific to political behavior. Finally, longitudinal studies could address the causal direction between liberal partisanship and the lower decision boundaries that we report in the present study. A more trivial potential explanation of the difference in RT between Liberal partisan and non-partisan groups observed here is that it was an artifact of task design, induced by interleaving party information and no party information conditions. This seems a less likely explanation given the observation that RT differences were evident even in the first few trials, with little change over the task. However, a variation of the same task with conditions varied by block would more thoroughly test this possibility. The eye tracking data supported the hypothesis that when only visual appearance was given as information, both groups employed decision processes analogous with those seen in economic choice: both groups exhibited significantly shorter final versus middle fixations as well as an increased likelihood of choosing the last seen option. K R A S T E V | 55 The choice behavior and eye movement findings in the no party information condition suggest that evidence drawn from appearance during voting choices is processed in a way similar to food choices studied in the past (Krajbich et al., 2010). These findings agree with other recent work showing that the aDDM can be applied to a broader range of tasks (Parnamets et al., 2014; Krajbich et al., 2012), and support the idea that the explanatory framework proposed for simple economic choices in this paradigm can be generalized to a variety of choices. Further work is needed to test the limits of this claim. As an initial step in exploring these limits, the second aim of this study was to examine the effects of partisanship on evidence gathering as indexed by choice behavior and eye movements. The party information condition adds further information, which we hypothesized would have a different influence on decision-making in Liberal partisans and non-partisans. Based on past research on the partisanship heuristic in low information elections (Schaffner & Streb, 2002), we expected that the former group would follow a simplifying heuristic, which would substantially alter decision behavior, leading to faster and more stereotyped (ie. ‘choose Liberal’) choices, insensitive to the social information from the faces that we assume guides decisions in the no party information condition. The choice probability and RT curves for partisans in the party information condition indicate that displaying a Liberal candidate during a trial makes partisans less sensitive to the ratings they previously gave based on a candidate’s appearance. In fact, there was no significant effect of rating on partisans’ choice probability or RT, suggesting that party information supersedes information drawn from the appearance of the candidates. However, the eye movement results tell a more nuanced story. K R A S T E V | 56 Final fixations were shorter than middle fixations in both groups across conditions. This result conforms to the predictions made by past studies of economic choice, supporting the idea that subjects may be engaging in evidence gathering until a decision threshold is reached. This finding does not allow us to identify whether partisans were using a “choose-my-party” heuristic or deliberating, as we would expect a barrier to be reached and thus interrupt the final fixation in both cases. However, we did predict that partisans in the party information condition would adopt a simpler search approach, quickly shifting their eyes to the red-framed candidate, resulting in fewer and shorter fixations. Although the difference in fixation duration only trended towards significance, partisans did indeed show significantly fewer fixations. Strikingly, this trend was also evident in partisans in the no party information condition. The last attended item data also support the conclusion that party information did not engage a heuristic that led to decisions entirely blind to any information except party affiliation. When the party affiliation of the last attended candidate was unknown and that candidate was previously rated 1 or 2 units higher than the alternative, partisans chose that candidate roughly 50% of the time. This supports the claim that partisans continue to gather evidence beyond party information, remaining at least somewhat sensitive to the factors that contributed to their prior value estimates based on candidate appearance as well. Although very little work has been done to explore how heuristics interact with evidence gathering, the findings here raise the possibility that party information does not engage a unique decision process, but rather integrates with other factors to drive a decision value toward a barrier. This possibility is supported by the little research done on the topic. In a study on branding, Philiastides and Ratcliff (2013) used a preference-based decision-making task and computational modelling to show that brand information and subjective preference are likely K R A S T E V | 57 integrated into a single source of evidence in the decision-making process. Further work is needed to clarify the mechanisms underlying decision-making heuristics, as well as to shed light on the neural substrates of these processes. Alternative Decision Models We have thus far framed this study in the context of past research on economic decision-making. However, our analysis makes assumptions about the nature of evidence gathering during choice that remain to be confirmed. Although models that describe integration of decision value toward one of two decision barriers such as the attentional Drift Diffusion Model have seen some success in predicting choice behavior (Ratcliff, 1978; Smith & Ratcliff, 2004), eye movements and brain activation patterns during economic choice, there is a larger set of viable models that describe evidence gathering during 2-Alternative Forced Choice tasks. Some of these, such as the Ornstein-Uhlenbeck model, are extensions of the formal logic behind the DDM and present a serial mode of evidence gathering. In the case of the OrnsteinUhlenbeck model, this is achieved by adding a “current accumulation state” term to the basic DDM equation which makes the current rate of evidence accumulation for each option dependent on the value accumulated for each option so far. Other models, classified as “accumulators”, rely on a parallel processing mechanism to gather evidence and are thus formally quite different from the DDM as they no longer represent a random walk. In one such popular alternative called the Race Model (Vickers, 1970), evidence is accumulated for both options in parallel (and is thus presumed to be unaffected by attention). The option which reaches a predetermined threshold first is the one which is ultimately chosen. In the Mutual Inhibition Model, on the other hand, evidence gathered in parallel for each alternative serves to dampen evidence gathering for its alternative (Usher & McClelland, 2001). K R A S T E V | 58 Although there has been significant support for both diffusion and accumulator type models in past years, the DDM (and aDDM variation) have provided better fits to the data from various perceptual and value-based choice tasks (Milosvljevic et al., 2010). Furthermore, electrophysiology studies have shown that the DDM may provide a plausible mechanism for visual discrimination at the neuronal level (Gold & Shadlen, 2007). Finally, the DDM is more parsimonious than many of the alternatives, as it uses fewer terms. This is an important feature of a model, as a higher number of parameters leads to more degrees of freedom and may thus cause over-fitting of the data. Nonetheless, since an important feature of a successful model is biological feasibility, future studies at the neural level will provide useful evidence for determining which model best describes value-based decision mechanisms in the brain. Conclusion & Future Directions This study explored a novel behavioral and conceptual framework that could inform research on the neural substrates of voting choices, as well as provide a basis for developing mathematical models of value-based choice that integrate beliefs gathered prior to the choice with online evidence gathering during the trial. There are uncertainties and limitations inherent to our design which should be addressed in future studies. As mentioned in the introduction, the binary choice task is a good starting point for comparing political choice to economic choice because it has been used widely in the past and provides the simplest possible choice. However, while binary choice is a good approximation of some classes of economic choices, it may not closely mimic the informationally complex choices involved in political behavior. In order to eventually reach an explanatory level that is useful in political science, future work must adapt the tasks used here in order to increase complexity. For example, future studies could start with the most basic version of the task reported here (binary choice modeled by K R A S T E V | 59 aDDM in non-partisans, with no party information) and increase complexity by adding other elements of interest such as political issues, a full range of political parties, known candidates and appearance ratings based on more factors. In this study, we have shown that participants exhibit similar choice behavior and eye movement patterns when making simple political choices to those observed in economic choice in the past. We have also shown that although party information strongly influences decision behavior in partisans, it does not do so in a way that completely supersedes other sources of information. Thus, models of evidence gathering that drive a decision value toward a barrier may generalize to more complex informational environments. The work presented here provides a starting point for studying political choice in the brain. Past work (Lim et al., 2011) has linked fixation-guided valuation to activity in the vmPFC and striatum, suggesting that these areas are responsible for encoding fixationdependent relative value signals. Future work using neuroimaging or lesion methods with similar task designs will be important in establishing the neural basis of political choice. K R A S T E V | 60 CHAPTER 4: GENERAL DISCUSSION & CONCLUSION This thesis made initial steps in investigating the conceptual, behavioral and neural intersections of economic and political decision-making. Progress toward understanding the decision processes behind political choice is not only relevant to political scientists, but is a step toward broadening the scope of decision neuroscience. Research in neuroeconomics provides a useful starting point for exploring the differences and commonalities between economic and political decision-making (Cacioppo & Visser, 2003). As a first step, here we sought to understand to what extent the fields of neuroeconomics and neuropolitics have taken comparable experimental approaches, and whether these have yielded similar results. We addressed this in Chapter 2 with a systematic review of the neuropolitics literature to date. We found that studies of political choice have been little influenced by the neuroeconomics literature, or the related and much broader animal literature on the brain basis of reward processing and value-based learning. Whereas studies of economic choice have had a strong experimental emphasis on tasks linking specific decisions to immediate reward outcomes, the same is not possible in a political choice context as the rewards of political choice are by nature ambiguous and time-delayed. Although none of the neuropolitics studies explicitly discussed reward, we examined the results of the subset of neuropolitics studies which used fMRI to report the neural correlates of positive and negative subjective value, looking for common patterns. We found very little commonality between the foci reported by neuropolitics studies and the regions of interest commonly related to subjective value in economic paradigms (Bartra at al., 2013). Although this result is limited by the small number of studies, it leads to two provisional conclusions: first, the neuropolitics literature to date has not taken advantage of frameworks developed to study economic choice in the brain; second, the neural correlates and perhaps K R A S T E V | 61 decision processes associated with political choice may be in part or entirely distinct from those observed in economic choice. Chapter 3 reports an effort to develop an experimental paradigm that bridges the study of economic and political choice. A binary choice task commonly used to study economic choice (Krajbich et al., 2010; Milosavljevic et al., 2010; Lim et al., 2011; Krajbich & Rangel, 2011; Krajbich et al., 2012) was adapted for the canonical political choice central to Western democracy: voting for political candidates. This allowed preliminary investigation of information gathering and deliberation during voting, as indexed by choice behavior and eye movements. Given that the fMRI meta-analysis reported in Chapter 2 did not yield overlapping regions of activation for economic and political choice, we aimed to establish whether choice behavior and eye movements followed similar patterns in experimental political choice as previously reported for ‘deliberative’ economic choice. We also explored whether these measures would reveal evidence of heuristic rather than deliberative processes under certain conditions, by studying the effects of partisanship in the presence and absence of party information. The deliberative-heuristic dichotomy is a central topic in the study of political behavior (Schaffner & Strebb, 2002). These two “types” of decision-making (assuming their conceptual distinction is legitimate) could potentially have distinct neural (Volz et al., 2006 & 2010), behavioral and eye movement correlates. Choice behavior and eye movement patterns from the first condition conformed to those seen in past studies of economic choice, providing preliminary evidence that deliberative political choice likely relies on a similar information gathering process as deliberative economic choice (Krajbich et al., 2010). Moreover, data showed that the partisan group made significantly fewer visual fixations, suggesting that, while they seemed to be engaging the same decision K R A S T E V | 62 processes as indexed by their overall patterns, they were systematically different in the amount of information needed to reach a decision. We addressed this formally with a preliminary modeling exercise, finding that the difference between the partisans and non-partisans could be approximated by a lowered decision value barrier (the amount of evidence in support of an option required before that option is chosen) within the aDDM. Choice behavior and eye movement pattern data from the second condition allowed us to test the effects of partisanship on political choice. Past work on political decision-making has characterized the partisanship heuristic as particularly prevalent in low-information elections, such as the one we created by providing nothing more than the visual appearance of candidates (Schaffner & Strebb, 2002). Thus, we expected party information to completely supersede visual appearance ratings in partisans. However, the data told a different story – although partisans were highly affected by party information, they exhibited the same choice behavior and eye movement patterns reported in economic paradigms thought to engage deliberative decisionmaking. Partisans took information conveyed by visual appearance into account and their choices were significantly affected by attention, as indexed by eye gaze. Although further work is needed to generalize these findings beyond Liberal partisans, these preliminary findings raise the possibility that “heuristic” choice differs in extent, rather than in kind, from deliberative choice. The behavior in both conditions, in both groups, could be accommodated by varying the parameters of the aDDM, for example. This is as a key future direction to pursue, of importance in both neuroeconomics and neuropolitics. In particular, our findings suggest that the framework used to study economic choice in the past, including but not limited to imaging studies, is applicable to political choice, as long as factors that are notably relevant in political choice, such as partisanship, are addressed in the design. K R A S T E V | 63 As studies of decision-making in the brain explore a growing number of ecologically valid components of choice (for example: risk (Knutson et al., 2008), uncertainty (Tobler et al., 2007), time-discounting (Kable & Glimcher, 2007), motivation (Bjork et al., 2010) and social interactions (Fliessbach et al., 2007)), they become increasingly relevant to the study of political choice, which incorporates many of these elements. However, if findings from such studies are to be used to grow our understanding of political choice, then future work must first characterize the exact role that these elements play in various kinds of political choice. The present thesis provides a framework that may be useful in understanding the roles of evidence gathering and heuristics in political choice. Furthermore, it attempts to lay the groundwork for a more systematic investigation of political choice that leverages the wealth of existing research and methodology present in neuroeconomics. Future work should aim to further bridge the gap between neuropolitics and neuroeconomics as both fields likely study decision behaviors that differ in degree rather than kind. These efforts have the potential to expand the explanatory limits of neuroeconomics, and are a valuable step toward gaining a deeper understanding of political choice which is, after all, a key driver of personal and social well-being. K R A S T E V | 64 ACKNOWLEDGEMENTS First and foremost, I would like to thank my supervisor Lesley Fellows for her continuous support, wisdom and patience in the creation of this thesis as well as in the ideation and execution of the associated experimental and conceptual work. Furthermore, I would like to thank the members of my advisory committee, Alain Dagher and Dietlind Stolle, as well as Elisabeth Gidengil, for their meaningful contribution throughout my degree. Next, I would like to express my gratitude to Avinash Vaidya for his constant aid at the lab and in the testing room and to Christine Déry for her help in recruiting research participants. In addition, I am thankful to past and present members of my lab (in alphabetical order, by last name) – Mathias Faeth, Ana Fernandez Cruz, Eldad Hochman, Anne Löffler, Alison Simioni, Ami Tsuchida and Chenjie Xia – for their assistance in my research and their feedback on presentations, conference materials, publication submissions and this thesis. Finally I would like to thank my mentor, Josephine Nalbantoglu, for her guidance. This research was supported by a grant from CIHR (MOP: 97821) and the Fonds de recherche du Québec-Société et culture. K R A S T E V | 65 REFERENCES Amodio D.M., Jost J.T., Master S.L., Yee C.M. (2007) “Neurocognitive correlates of liberalism and conservatism.” Nature Neuroscience 10: 1246 – 1247. Badre D., Poldrack R.A., Pare-Blagoev E.J., Insler R.Z., Wagner A.D. (2005) “Dissociable controlled retrieval and generalized selection mechanisms in ventrolateral prefrontal cortex.” Neuron. 47: 907–918. 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